Geographical restriction to refuges implies the regional extinction of taxa in areas of the previous range falling outside the refuge. A comparison of the circumstances in the refuge with those in areas from which the taxa were eliminated is potentially informative for pinpointing the causes of extinction. A synthesis of data on the geographical and stratigraphical distributions of cool-water molluscs of the North Pacific and North Atlantic Oceans during the late Neogene reveals four patterns of geographical restriction, at least two of which imply that climatic cooling was not the only cause of extinction during the last several million years. These four patterns are (1) the northwestern Pacific restriction, involving 15 taxa whose amphi-Pacific distributions during the late Neogene became subsequently restricted to the Asian side of the Pacific; (2) the northwestern Atlantic restriction, involving six taxa whose early Pleistocene distribution is inferred to have been amphi-Atlantic, but whose present-day and late Pleistocene ranges are confined to the northwestern Atlantic; (3) a vicariant Pacific pattern, in which many ancestral amphi-Pacific taxa gave rise to separate eastern and western descendants; and (4) the circumboreal restriction, involving six taxa whose early Pleistocene distribution, encompassing both the Atlantic and Pacific Oceans, became subsequently limited to the North Pacific. Like the Pliocene extinctions in the Atlantic, previously studied by Stanley and others, the vicariant Pacific pattern is most reasonably interpreted as having resulted from regional extinction of northern populations in response to cooling. The northwestern Pacific and Atlantic restrictions, however, cannot be accounted for in this way. In contrast to the northeastern margins of the Pacific and Atlantic, the northwestern margins are today characterized by wide temperature fluctuations and by extensive development of shore ice in winter. Northeastern, rather than northwestern, restriction would be expected if cooling were the overriding cause of regional extinction. Among the other possible causes of extinction, only a decrease in primary productivity can account for the observed northwestern and circumboreal patterns of restriction. Geographical patterns of body size and the distribution of siliceous deposits provide supporting evidence that primary productivity declined after the Miocene in the northeastern Pacific, but remained high in the northwestern Pacific, and that productivity in the Pacific is generally higher than it is in the Atlantic. The patterns of geographical restriction in the northern oceans thus provide additional support to previous inferences that reductions in primary productivity have played a significant role in marine extinctions.
Phylogeography and connectivity of thePseudocalanus(Copepoda: Calanoida) species complex in the eastern North Pacific and the Pacific Arctic Region